ACT A PAL A EON T 0 LOG IC A POLONICA Vol. XVI 1971 No. (

CYPRIAN KULICKI

NEW OBSERVATIONS ON RHABDOPLEURA KOZLOWSKII () FROM THE BATHONIAN OF POLAND

Abstract. - Specimens of Rh. kozlowskii Kulicki, 1969 have here been described from the Bathonian of southern Poland. A secondary layer, never observed before, has been found inside zooidal tubes.

INTRODUCTION

At present, the genus Rhabdopleura is represented by at least two species clearly different from each other, Le. Rh. normani Allman, 1869 and Rh. striata Schepotieff, 1909. All other Recent species display a con­ siderable similarity to Rh. normani and the necessity to distinguish them is called in question by many investigators (Schepotieff, 1906; Dawydoff 1948; Thomas & .Davis, 1949; Burdon-Jones, 1954 and others). The species Rh. compacta Hincks, 1880 has recently been restored by Stebbing (1970), who concludes that Rh. compacta differs from Rh. normani mostly in the form of colonies and lack of ring-shaped part of the stolon. The following three fossil species have hitherto been described: Rh. vistulae Kozlowski, 1956 from the Danian of Poland, Rh. eocenica Tho­ mas & Davis, 1949 from the of England, and Rh. kozlowskii Kulicki, 1969 from the Callovian of Poland. The specimens of Rh. kozlowskii, described in the present paper, were etched with hydrochloric acid fTom ca1careous-marlyconcretions which occur in black and dark-gray Bathonian clays, Morrisiceras morrisi Zone (R6zycki, 1953) of Blanowice near Zawiercie. The concretions, varying in shape, are mostly spherical or ellipsoidal and fluctuate in size between a few and some scores of centimetres. Many of the concretions collected contain a macrofauna of molluscs or pieces of wood. The remains of hydroids, scolecodonts, organic linings of foraminifers, plant remains and other, unidentified remains, including many similar to the Callovian ones 416 CYPRIAN KULICKI from Luk6w (Kulicki, 1969), were etched out along with the remains of Rhabdopleura. The specimens of Rh. kozlowskii from the Bathonian of Blanowice are preserved much the same as those fram the Callovian of I:.uk6w and are represented by these same elements of tubarium. Specim€ns of Rh. normani from Bergen, Norway, have been used by the present writer as a comparative material. The terminology of the elements of tubarium has been adopted according to Kozlowski (1956) and the writer's previous work (Kulicki, 1969). Schepotieff's definition "st€ril Knospe" (sterile bud) was replaced by the writer (after Stebbing) with the term "dormant bud". Schepotieff (1907) believed that the buds contained in cysts, could not be turned into zooids. Stebbing studies (1970) have, however, shown that such buds may develop into normal individuals and, therefore, the term a dormant bud is more appropriate for them. A diagram of fundamental measurements of various elements of tubarium is shown in Fig. 1.

A

m

B

Fig. 1. - Diagram of fundamental measurements of zooidal (A) and stolonal (B) tubes of the genus Rhabdopleura: 1 inner diameter, 2 outer diameter without fusellar collars, 3 outer diameter with fusellar collar, 4 width of fusellar collar 5 thicknes·s of wall of zooidal tube, 6, 7 width of fusellus, 8 outer diameter of stolonal tube, m marginal membrane.

The specimens of Rh. kozlowskii and Rh. normani described in the present paper are housed at the Palaeozoological Institute, Polish Academy of Sciences, for which the abbreviation Z. Pal. is used. RHABDOPLEURA KOZLOWSKII FROM THE BATHONIAN 417

DESCRIPTION

Family Rhabdopleuridae Harmer, 1905 Genus Rhabdopleura Allman, 1869 Rhabdopleura kozlowskii Kulicki, 1969 (Text-figs. 2-7)

1969. RhabdopLeura kozLowskii Kulicki, 1969; C. Kulicki, The discovery..., pp. 540, 542, 544, 545; Figs. IE, 3B, 4A, 5J.

Material. - Thirty-one fragmentary zooidal tubes, seven fragmentary stolonal tubes, 23 variously-sized fragments of stolons and 69 cysts of dormant buds, some of them with stolons. Description. - Zooidal tubes. A diameter of the erect parts of zooidal tubes, the width of fusellar collars not taken into account, varies within limits of 124 and 189 fl, averaging 154 1ft, whereas a diameter of tubes including fusellar collars amounts to 143-235 fl, averaging 178 f-l; a mean width of a fusellar collar amounts to 12 fl. The width of fuselli varies from 20 to 53 !-l, averaging 42/l. Within the range of particular zooidal tubes, the width of fuselli is considerably variable although the differences do not exceed extreme values. In many specimens from Blanowice and Luk6w, one or two layers of skeletal substance, similar to that which forms the fuseliar part of tube and which is termed by the writer a secondary layer (Figs. 3, 7) occur inside zooidal tubes under the fuselar layer. The secondary layer does not display any traces of fusella'r structure. All injuries and cracks of the secondary layer (d. Fig. 3) run along the longitudinal axis of tube. A peculiar texture of secondary layer with parallel and closely spaced, dark striae running along the longitudinal axis of tube are visible in strong magnification. No such texture can, on the other hand, be observed in the fusellar layer. The secondary layer IS frequently thicker inside tubes with small than normal and large diameters. In one case, the secondary layer is double (Fig. 3) and in many specimens it is not observed at alL Stolonal tubes. The width of stolonal tubes, minus the marginal membrane, amounts to 130 to 158 !-t, averaging 154 !-t, and the width of their fuselli fluctuates within limits of 30 and 58, averaging 39 !-t. The lower, flat wall of stolonal tubes is frequently lacking. It is much thinner than the convex wall. Stolons. In transverse section, stolons are semicircular, with an upper wall convex and lower flat and several times thinner than the convex one. The width of stolons varies from 25 to 51 !-t, averaging 38 !-t. Such a variability is even observed within a specimen of a bifurcated stolon. In places of bifurcation, a lateral stolon may be separated from the main B

D E

Fig. 2 RHABDOPLEURA KOZLOWSKII FROM THE BATHONIAN 419 one by a few diaphragms having 7-9 f..L wide pores. The diaphragms are mostly situated in contractions of the vesicular extensions of the proximal part of the lateral stolon. Peduncular and lateral stolons detach them­ selves alternately from the main one at intervals of about 310 to 630 f..L. Peduncular stolons are short and vesicular, mostly provided with two or three diaphragms having 4-9 f..L wide (averaging 7 f..L) pores. Peduncular stolons with a higher number of diaphragms were also observed (Fig. 5

B, C3).

c

2.

A B

Fig. 3. - Rhabdopleura kozlowskii Kuticki (Z. Pal. Pb. II/8-10): A a zooidal tube, Bathonian, Blanowice; B-C zooidal tubes, Callovian, Luk6w: 1 a fusellar layer of a zooidal tube, 2a, 2b particular laminae of a secondary layer.

Cysts of dormant buds. The cysts of dormant buds resemble in shape the proximal part of a contractile stalk of a zooids (Schepotieff, 1907) and they are formed on its basis (Stebbing, 1970). The width of the cysts of dormant buds, which closely adhere to the waUs of stolonal tubes, varies

---- ~~------Fig. 2. - A Rhabdopleura sp., Bathonian, Blanowice: zooidal tubes connected by common fuselli (Z. Pal. Pb. II/I); B-F Rh. kozlowskii Kulicki, Bathonian, Blanowice: erect part of zooidal tubes (Z. Pal. Pb. II/2-7); G-H Rh. normani Allman, Recent, Bergen, Norway: bifurcated zooidal tubes (Z. Pal. Pb. 1/101). A

E

F

Fig. 4 RHABDOPLEURA KOZLOWSKII FROM THE BATHONIAN 421 within limits of 85 and 166 11, averaging 129f!, and their length 260 and 460 !!, averaging 344 !!. Remarks. - The dimensions of the specimens of Rh. kozlowskii from the Bathonian of Blanowice are contained within the range of individual variability of the specimens of this same species from the Callovian of Luk6w. Mean dimensions of the elements of tubarium are approaching each other in the two forms. In other respects, the stolonal tubes and cysts of dormant buds in the Bathonian form do not differ radically from the Callovian one or from the remaining species, except for Rh. striata. A certain number of peduncular stolons, divided by a larger number of diaphragms and being longer than those of the Callovian, specimens from Luk6w (Fig. 5, B, C3), which occur among the Bathonian specimens make up a small differences between them. This difference is, however, too small to constitute a basis for the distinction of the Bathonian form as a separate species. As follows from Kozlowski's (1956) and Kulicki's (1969) studies, the structure of peduncular stolons and the shape of the transverse section of the main stolon are of a decisive importance to distinguishing species. The secondary layer, internally lining the ascending parts of zooidal tubes, is a quite new skeletal element observed in Rh. kozlowskii only. The occurrence of a thicker or bilaminate secondary layer inside the tubes with small or normal diameters or its frequent lack in the tubes with large diameters, indu-ce the present writer to conclude that the secondary layer played a reinforcing role. Its presence and degree of thickening depend on the length and diameter of the tubes and on envi'ronmental conditions. What is known as longitudinal ribs, occurring inside woidal tubes between the creeping and ascending part of a tube was described by Schepotieff '(1907) in Rh. normani as reinforcing elements. This author found that, in the case of short zooidal tubes, the longitudinal ribs do not occur at all or are poorly developed. No ·reinforcing elements have been found in Rh. vistulae Kozlowski. Figs. 2 Al and A z show zooidal tubes, which closely adhere to each other and whose fusellar systems are connected with each other by common fuselli. This close connection might be formed in the case of a simultaneous growth of both tubes, which probably were parts of one and the same colony. Due to their small diameters (without fusellar collars amounting to 90 and 133 l.L and with fusellar collars - to 101

Fig. 4. - Rhabdopteura koztowskii Kulicki, Bathonian, Blanowice (Z. Pal. Pb. II/1l-16): A a stolonal tube with a transverse septum, dorsal view; BI- 3 a fragment of a colony, BI dorsal view, Bz ventral view, B3 lateral view; C-D stolonal tubes, ventral view: E-F zooidal tubes, qs transverse septum, ct creeping part of a zoo­ idal tube, et erect part of a zooidal tube, st stolonal tube, p attachment place of a zooidal tube. 422 CYPRIAN KULICKI

and 143 J..L) and width of fuselli, smaller than in the remammg specimens from Luk6w and Blanowice (24 and 25 1-1), their assignment to Rh. kozlow­ skii and other known species remains uncertain.

At

Cz Cl

A

C4 C C3

O,5mm B Fig. 5. - Rhabdopleura kozlowskii Kulicki, Bathonian, Blanowice (Z. Pal. Pb. II/17-19): A-B bifurcated stolons with peduncular stolons; C a bifurcated stolon with five, clearly visible peduncular stolons, C1-C2 peduncular stolons with two diaphragms, C3 a peduncular stolon with at least four complete and one incomplete diaphragm C4 a peduncular, abnormally developed stolon. RHABDOPLEURA KOZLOWSKII FROM THE BATHONIAN 423

In Rh. kozlowskii, the proximal parts of ascending zooidal tubes have smaller diameters than the distal parts. Besides, they have poorly de­ veloped fusellar collars and, frequently, narrower fuselli (Figs. 2E, 4B). A similar growth of zooidal tubes, which may be explained by a zooid growth in the process of forming the tube may sometimes be observed

A2 A1

F E

11 12 J 2

O,5mm K

Fig. 6. - A-B RhabdopLeura normani Allman, Recent, Bergen, Norway; cysts of dormant buds (Z. Pal. Ph. 1/101): Al ventral view, A 2, B dorsal view; C-K Rh. kozLowskii Kulicki, Bathonian, Blanowice; cysts of dormant buds (Z. Pal. Ph. II/20- 28): C2 , Dl , E-H, It, J l ventral view, Cl , D 2, 12, h dorsal view.

8 Acta Pal..eontologica Polonica nr 4/71 424 CYPRIAN KULICKI in Rh. normani (Schepotieff, 1907). Bifurcated zooidal tubes were observed by Kozlowski (1948, Fig. 14) and the present writer (Figs. 2 G, H) in the Recent material. In such cases, one of the branches is always older than the other. Impurities of various types occur in the distal parts of older tubes. Most likely, these impurities got inside 'the tubes in the process of and after the degeneration of an zooid when it was incapable of removing them. In the case of the regeneration of this same zooid, it used the same zooidal tube, if it was not excessively filled with impurities or, if it was, only its proximal part, with a simultaneous construction of a new zooidal tube, which was built after perforating an opening in the old tube. Such an explanation results from the facts that the new tube beings with a smaller diameter and narrower fuselli and has less prominent fusellar collars than its distal part, which allow one to suppose that a re­ peated growth of the zooid took place as a new tube was formed. The processes of degeneration and regeneration of the zooids were described by Schepotieff (1907) and Stebbing (1970). Ecological Temarks. - Recent representatives of Rhabdopleura are widely distributed over the Earth. In the southern hemisphere, they were recorded near Celebes, New Zealand, Tasmania, South Australian coasts and Antarctica (Johnston, 1937). In the northern hemisphere, they occur near Ceylon (Schepotieff, 1909) in they Bay of Marseilles, in the Atlantic Ocean, in the Bay of Biscay, in the North Sea, Norwegian Sea, Barents Sea, in Davis Straits near Greenland, as well as in the Arctic Sea near Spitsbergen (d. Burdon-Jones, 1954 and Stebbing, 1970). The depths at which the genus occurs range from 2 m in the case of Rh. striata (Sche­ potieff, 1909) and 5 m of Rh. normani (Burdon-Jones, 1957) to 896 m (Stebbing, 1970). The salinity of waters in which Rh. normani is found fluctuates within limits of 33 and 35%0 (Burdon-Jones, 1954) and their temperature -1°C (Stebbing, 1970) and + 12°C (Burdon-Jones, 1954). Recent colonies of Rhabdopleura occur attached on other organisms such as corals, bryozoans, serpulids or on the shells of mollUSCS, on stones and even directly on a rocky, stony, gravelly, sandy, clayey and loamy bottom. They are accompanied by molluscs, tunicates, brachiopods, bryo­ zoans, serpulids and other polychaetes, corals, hydroids, sponges and foraminifers (Burdon-Jones, I.e., Stebbing, I.e.). Rh. kozlowskii Kulicki, 1969 found at Luk6w occurs in calcareous concretions embedded in clays. These same concretions contain the re­ mains of echinoderms, shells of molluscs, brachiopods, bryozoans, poly­ chaetes, hydroids, spicules of sponges, foraminifers and remains of land plants (Makowski, 1952; Kulicki, 1969). Likewise, specimens of Rh. koz­ lowskii from the Bathonian of Blanowice were found in calca~eous-marly concretions occurring in black and gray clays, together with molluscs, hydroids, polychaetes, foraminifers and remains of land plants. The genera RHABDOPLEURA KOZLOWSKII FROM THE BATHONIAN 425

III II I(

If

II'I I, ~II (I 1\, ~I

II II III II

A B C D

Fig. 7. - A-C Rhabdopleura kozlowskii Kulicki, Callovian, Luk6w: longitudinal section through the wall of zooidal tube in its erect part; a secondary layer variable in thickness clearly visible under a fusellar layer (Z. Pal. Pb. 11/29-31); D Rh. normani Allman, Recent, Bergen, Norway: a longitudinal section through the wall of a zooidal tube in its erect part (Z. Pal. Pb. IIIOl).

of gastropods from the Callovian of Luk6w: Turbo, Trochus, Natica, Tur­ ritella and Ceritium, as well as pelecypods: Lima, Pinna and Leda occur· at present from the water level down to a depth of 400 m (Davitashvili & Merklin, 1966, 1968). A mean depth of their occurrence amounts to about 135 m. The Recent Rh. normani most frequently occurs at a depth between 100 and 300 m (Schepotieff, 1906). Since the concentration of fauna in

8' 426 CYPRIAN KULICKI the Luk6w concretions was probably accompanied by waving (Makowski, 1952, p. IX), we may assume that the depth of a sea basin, in which the Callovian fauna lived, was approaching 100 to 150 m and, therefore, it was contained within the range of the optimum occurrence of Rh. nor­ mani. Likewise, the temperature requirements of Rh. kozlowskii are within limits characteristic of a Recent species. On the basis of ammonites from Luk6w and using the isotopic methods, Stahl & Jordan {1969), have evaluated a palaeotemperature at 12.5-13.5°C, which may be placed near the upper limit of Rh. normani's temperature requirements. The type of deposits and faunal associations in which Rh. eocenica and Rh. vistulae occur are similar to those of Rh. normani and Rh. kozlowskii. It is also the Eorhabdopleura urbaneki Kozlowski, 1970, displaying a great morphological similarity to the genus Rhabdopleura Allman (Kozlowski, 1970) 1, that probably lived at moderate depths (found in an organodetrital limestone). Considering that all fossil species of the genus Rhabdopleura were found in either neritic, or littoral deposits to­ gether with shallow water fauna, we may state that the Rhabdopleuridae did not change their ecological requirements, at least from the and even Ordovician. This is in conformity with Kozlowski's (1970) state­ ment that the genus Rhabdopleura makes up a tribe strongly conservative both morphologically and ecologically.

Palaeozoological Institute of the Polish Academy of Sciences Warszawa, Zwirki i Wigury, 93 September, 1970

REFERENCES

BURDON-JONES, C. 1954. The habitat and distribution of Rhabdopleura normani Allman. - Naturvitensk. Rekke, 11, 3-17, Bergen. DAVITASVILI, L. & MERKLIN, R. 1966. Spravocnik po ekologii morskich dvu­ stvorok. 5-349, Moskva. & 1968. Spravocnik po ekologii morskich briuchonogich. 5--169, Moskva. DAVYDOFF, C. 1948. Class des Pterobranches. In: P. Grasse, Traite de Zoologie, 2, 454-489, Paris. JOHNSTON, T. H. 1937. Australian Antarctic Expedition 1911-1914. - Sci. Reports, C, Zoo1. Botany, 3, 4, 2-8, Sydney. KOZLOWSKI, R. 1948. Les Graptolithes et quelques nouveaux groupes d'animaux du Tremadoc de la Pologne (Graptolity i par~ nowych grup zwierzqt z tre­ madoku Polski). - Palaeont. PoL, 3, I-XII+1-235, Warszawa.

1 A tentative assignment of this species to a new genus results from the structure of its stolons being unknown, although the known elements of tubarium display a great similarity to these same elements in other representatives of the genus Rhabdopleura Allman (Kozlowski, 1970). RHABDOPLEURA KOZLOWSKI! FROM THE BATHONIAN 427

1956. Sur Rhabdopleura du Danien de Pologne (Rhabdopleura z danu Pol­ ski). - Acta Palaeont. Pot., 1, 1, 12-21, Warszav.ra. 1970. Nouvelles obs'ervatiQns sur les Rhabdopleurides (Pterobranches) Ordo­ viciens (Nowe obserwacje nad ordQwickimi Rhabdopleurida (Pterobranchia) ­ Ibidem, 15, 1, 3-17. KULJCKI, C. 1969. The discovery of Rhabdopleura (Pterobranchia) in the Jurassic of Poland (Odkrycie Rhabdopleura (Pterobranchia) w jurze Polski). - Ibidem, 14, 4, 537-55l. MAKOWSKI, H. 1952. La Faune Callovienne de Luk6w en Pologne (Fauna kelowej­ ska z Lukowa). - Palaeont. Pol., 4, I-X+1-64, Warszawa. ROZYCKI, S. Z. 1953. G6rny dogger i dolny maIm Jury Krakowsko-Cz~stochow­ skiej. - Prace Inst. Geot., 11, 110-115, Warszawa. SCHEPOTIEFF, A. 1906-1907. Die Pterobranchier. - Zoot. Jb. Anat. etc. (2), 23/24, 463-534, 193-238, Jena. - 1909. Die pterobranchier des Indischen Ozeans. - Zool. Jb. Syst. etc. (1), 28, 429-448, Jena. STAHL, W. & JORDAN, R. 1969. General considerations on isotopic palaeotempe­ rature determinations and analyses on Jurassic ammonites. - Earth Planet. Sci. Lett., 6, 173-178, Amsterdam. STEBBING, A. R. D. 1970. The Status and ecology of (Hemichordata) from Plymouth. - J. Mar. Biol. Assoc. Un. Kingdom, 50, 1, 209-221, Cambridge. 1970. Aspects and life cycle of Rhabdopleura compacta (Hemichordata). ­ Int. J. Life Oc. Coast. Waters, 5, 3, 205-212, Heidelberg, New York. THOMAS, H. D. & DAVIS A. G. 1949. The pterobranch Rhabdopleura in English Eocene. - Bult. Brit. Muss. (Nat. Rist.), Geol., 1, 1, 1-19, London.

CYPRIAN KULICKI

NOWE OBSERWACJE NAD RRABDOPLEURA KOZLOWSKII (PTEROBRANCHIA), Z BATONU POLSKI

Streszczenie

W pracy tej opisano okazy Rh. kozlowskii Kulicki, 1969 znalezione w batotiskich konkrecjach w gliniance w Blanowicach kolo Zawiercia. Gatunek ten znany byl dotychczas z keloweju Lukowa. Nowe znalezisko obnizylo do. batonu dolnq granic~ geologicznq wYlSt~powania rodzaju. W por6wnaniach formy batotiskiej z kelowejskq nie stwierdzono wi~kszych r6znic ilosciowych i jakosciowych. Opisano nowy ele­ ment wzmacniajqcy, rozwini~ty we wznoszqcej si~ cz~sci rurek zoidalnych w posta­ ci warstv,;y wt6rnej, nie wykazujqcej budowy fuzelarnej, wyscielajqcej wewn~trznq powierzchni~ rurek. Znane u wsp6lczesnego gatunku Rh. normani Allman wewn~trz­ ne zeberka podluzne pelnily zapewne podobnq rol~, lecz mniej wszechstronnie niz 428 CYPRIAN KULICKI warstwa wt6rna. Autor uwaza, ze obecnosc warstwy wt6rnej uzalezniona jest od srednicy i dlugosci rurek zoidalnych, a takze od wplyw6w srodowiska zewnt:trznego. W wyniku por6wnan morfologicznych i ekologicznych, autor potwierdza wniosek Kozlowskiego (1970), ze rodzaj Rhabdopleura jest szczepem bardzo starym i kon­ serwatywnym pod wzglt:dem morfologicznym i ekologicznym.

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Pe3lO.Me

B pa60Te on~caHbI 3K3eMnJmpbI Rh. kozLowskii Kulicki, 1969, HaH~eHHble B 6aTc­ K~X KOHKpe~~HX B rJI~HHHOM Kapbepe MeCTHOCT~ BJIHHOBJ1~e 6JIJ13 r. 3aBep~e. 3TOT BJ1~ ~o CJ1X nop 6bIJI J13BeCTeH B KeJIJIOBee paHoHa r. JIyKyB. HOBoe MeCTOHaXOJK~eHJ1e nepe~BJ1HYJIO B 6aTcK~H HPYC HJ1JKHIOIO rpaHJ1~y pacnpOCTpaHeHJ1H po~a. IIpJ1 cpaB­ HeHJ1J1 6aTcKoH J1 KeJIJIOBeHCKOW lPOpM CYll.\eCTBeHHbIe pa3JIJ1'iJ1H KOJIJ1'ieCTBeHHoro J1 Ka'ieCTBeHHoro xapaKTepa He 6bIJIJ1 BbIHBJIeHbI. OnJ1CaH HOBbIW YKperrJIHIOll.\~H 3JIeMeHT B BOCXO~Hll.\eH 'iaCTJ1 300J1~HbIX Tpy60K B BJ1~e BTOpJ1'iHOrO CJIOH, He rrpo­ HBJIHIOll.\erO lPI03eJIJIHpHOH CTpyKTypbI, BbICTJ1JIaIOll.\erO BHyTpeHHIOIO rrOBepXHOCTb Tpy60K. li3BeCTHhle y COBpeMeHHoro BJ1~a Rh. normani Allman BHyTpeHHJ1e rrpO~OJIh­ HhIe pe6pa ~rpaIOT, BepoHTHo, nO~06HYIO pOJIh, O~HaKO He HaCTOJIbKO BceCTOpoHHIOIO KaK BTOPJ1'iHhIW CJIOW. ABTOP rrpe,IJ.nOJIaraeT, 'iTO HaJI~'iJ1e BTOpJ1'iHOrO CJIOH 3aBJ1CJ1T OT ,IJ.J1aMeTpa J1 ,IJ.JIJ1HhI 300~,IJ.HhIX Tpy60K, a TaKJKe OT YCJIOBJ1H BHeWHeH cpe~bI. Ha OCHOBaHJ1J1 MOPlPOJIOr~'ieCKJ1X J1 9KOJIOrJ1'ieCKJ1X conOCTaBJIeHJ1W aBTOp rro~~epJKJ1BaeT B3rJIHA K03JIOBCKOro (1970), KOTOpbIW rJIaCJ1T, 'iTO POA Rhabdopleura HBJIHeTCH ,IJ.peB­ HJ1M J1 KOHCepBaTJ1BHbIM TJ1nOM B MOPlPOJIOr~'ieCKOM J1 3KOJIOrJ1'ieCKOM OTHOlIIeH~HX.